Spatial distribution,population dynamics and productivity of <Emphasis Type="Italic">Spisula subtruncata</Emphasis>: implications for <Emphasis Type="Italic">Spisula</Emphasis> fisheries in seaduck wintering areas

Bivalves are important in shallow marine habitats, not at least being the major food resource for seaducks such as the common scoter (Melanitta nigra), thousands of which are wintering on the Western Coastal Banks, near the Belgian-French border (North Sea). Next to this ecological importance, fishable stocks of one of these bivalves, Spisula subtruncata, occur in the area. This study aimed at investigating S. subtruncata’s spatial distribution, population dynamics and productivity and its implications for a sustainable Spisula fishery in seaduck wintering areas. The spatial distribution of S. subtruncata was studied in 1994 and 1997 in two areas of the Belgian Western Coastal Banks. The population dynamics and production were investigated by monthly sampling of two stations between April 1995 and April 1996 and a seasonal sampling between April 1996 and April 1998. Spisula subtruncata had a patchy distribution in the deeper (6 m), fine sandy (200 ± 20 μm) sediments of the Abra alba community, mainly found in the western most part of the Western Coastal Banks. In August 1995, an overwhelming and successful recruitment was observed in this area: local densities were as high as 150,000 ind m⁻². Minor, non-successful recruitments were detected in August 1996 and 1997. Due to space limitation, high densities of S. subtruncata are hypothesized to be responsible for the occurrence of aberrant shapes as observed from August 1996 onwards. Growth was described by a seasonally oscillating version of the von Bertalanffy growth function (VBGF): a growth stop was observed from late autumn till early spring. The VBGF parameters K (growth constant) and L _∞ (asymptotic length) were estimated at 0.7–0.9 and 31–33 mm. A combination of length and individual biomass increment showed: (1) a faster length increment of smaller individuals during the second growing period (catching-up phenomenon), (2) a constant length combined with a decreasing individual biomass during the suboptimal winter periods (except for the first winter, when the individual biomass slightly increased), (3) a positive relationship between the individual biomass decrease and the seawater temperature during the winter periods, and (4) a strong increase of the individual biomass in early spring (April 1997 and 1998) because of gametogenesis, followed by a decrease because of spawning (August 1997). The extremely high total production of the 1995 year class in the tidal gully (Potje) during the study period was estimated at approximately 1,500 g ash-free dry weight (ADW) m⁻² or 600 g ADW m⁻² on average per year. Shellfisheries for S. subtruncata within seaduck wintering areas, such as the Western Coastal Banks, should be carefully deliberated since (1) an important food resource for the seaducks will decrease, (2) the ecologically most diverse and rich macrobenthic A. alba community will be heavily affected, and (3) the recovery of Spisula populations after depletion is expected to be erratic.     

Life cycle, seasonal vertical distribution and feeding of Calanus finmarchicus in Skagerrak coastal water

 Day/night zooplankton sampling in Kosterfjorden, Sweden, gave information on population succession, vertical distribution and feeding of Calanus finmarchicus over 17 months. Copepodid Stage 1 and 2 (C-I, C-II) were present from December to August, indicating reproduction during most of the year. Mating and breeding for the overwintering generation mainly took place in February/March, resulting in peak abundance of C-I in March/April. Secondary breeding periods were in April and July/August, but the resulting recruitment from these were low. The relative recruitment success in the first spring was around 13 times higher than during the second spring. Low temperature in the surface water early in the year and depression of the spring phytoplankton bloom may have caused the failure in the second year. Population biomass peaked at >15 g dry weight m⁻² during the numeric peak of the youngest stages in March/April. A more sustainable level of high biomass of 8 to 10 g dw m⁻² was gradually built up during summer, mainly due to a continuous accumulation of C-Vs. Adults and C-Vs comprised the overwintering population, with 7 to 14% and 85 to 93%, respectively, for the 2 years, but only C-Vs staying in the deep water were in a resting state. Adult males showed a strong diurnal vertical migration (DVM) of the usual type from spring to early autumn and a reversed DVM during the cold season. They were in a feeding state throughout the year. Adult females showed the usual type of DVM during summer to autumn, but commonly a reversed DVM during winter to spring. They were usually in a feeding state, with no pronounced differences between surface and deep water or between day and night. C-Vs aggregated in the deep water from October to March and performed DVM in April to June. They were commonly feeding in the deep water between March and June but showed no or very low feeding activity there from July to February. C-Vs in the surface water were commonly feeding and showed the highest proportion of feeding in autumn, when the population in the deep water was inactive. C-V constituted up to nearly 100% of the population biomass, and therefore must be of profound ecological importance. Defined by this dominant role, the population of C. finmarchicus can be characterised as having an active period of feeding, reproduction and development from February to July with a following 6 to 7 months of resting in the deep water, when development is arrested and no feeding occurs. Received: 1 October 1999 / Accepted: 27 April 2000     

Growth,stomach contents and parasites in a tidepool population of the hippolytid shrimp Heptacarpus futilirostris,with reference to reproduction

The biology of the hippolytid shrimp Heptacarpus futilirostris (Bate) was studied in a tidepool at Kominato, central Japan from January 1983 to June 1984. In males, the relative growth ratio between the third maxilliped length and body length revealed a turning point at approximately 15 mm in body length. Population recruitment was presumed to continue from April to December. The range of body length did not increase from July to October, and thereafter increased rapidly in both sexes up to a maximum range, i.e. from 6 to 29 mm in January. The wide range was presumably due to the simultaneous occurrence of rapid growth with population recruitment. The growth rates of size groups above and below 15 mm were different in the males observed in January. Large males disappeared, probably due to death, and small males participated in reproduction after July. The growth rate increased from late autumn to spring and decreased from summer to early autumn, and was influenced by relative food abundance, especially seaweeds. Two parasitic isopods were identified: Bopyrinella antilensis nipponica in the branchial cavity, and Epiphryxus sp. on the abdomen. It is suggested that parasites have more influence on maturity than on growth. Ovigerous females were found from January to October, with the breeding season peak between April and June. The relationship between female body length (L) and clutch size (N) was expressed by the regression equation: N=5.11 L^1.6347 (r=0.57). This species is identified as a multiple breeder. The relatively large number of larvae presumably compensates for the low larval survival rate.     

Dynamics and biological cycle of a Melinna palmata (Ampharetidae) population during the recolonisation of a dredged area in the vicinity of the harbour of Brest (France)

The reproductive cycle and growth parameters were studied in a Melinna palmata population, for 3 yr after dredging, in the vicinity of the harbour of Brest, France. This species first settled in May 1979, 8 mo after dredging had ceased. Measurements of the diameter of coelomic oocytes indicated that spawning took place from August to October and settlement in autumn and early winter. The first settled cohort displayed a high growth rate (0.25 mm width mo^-1 in summer) and a low longevity (22 mo); the growth rate of the second settled cohort was not significantly different from the first, but its longevity was higher (28 mo); the third settled cohort had a lower growth rate (0.15 mm width per month), but lower mortality and therefore a longer life expectancy. A fourth cohort appeared in August 1981 but was not followed because of the termination of the study. M. palmata colonised the dredged area in two ways: by settlement of juveniles in autumn and by probable immigration of young and adults in May. The population increased in number and biomass from the end of dredging (August, 1978) to June 1981 and decreased after this period. The demographic strategy of this species changed from r- to K-selection during this recolonization as a result of intraspecific and interspecific competition.     

Biology,population dynamics and secondary production of<Emphasis Type="Italic"> Tylos europaeus</Emphasis> (Isopoda,Tylidae) on the western coast of Portugal

The biology, population dynamics, and production of Tylos europaeus were studied in two sandy beaches of the western coast of Portugal. At both sites, reproduction occurred seasonally, from April to July, with only one new cohort produced per year. Regarding population dynamics, cohort-splitting events were detected in males at the beginning of the reproduction period (April/May), resulting in two groups with distinct growth rates (fast-growing vs slow-growing males). Different biological characteristics were consequently detected in these two groups, namely regarding body size, lifespan, and contribution to the reproductive effort. Lifespan was estimated as approximately 3 years, for females and fast-growing males, and 4 years for slow-growing males. Cohort-splitting among males appeared as a possible strategy to cope with the highly male-biased sex ratios observed, which could lead to a strong male-male competition for mating. T. europaeus appeared as an annual species, with a univoltine life-cycle (one generation per year), and iteroparous females reproducing twice during their lifespan. Average growth production (P) was estimated at 0.082 g.m^–2.yr^–1 AFDW (ash-free dry weight) and the average annual biomass () (standing stock) at 0.052 g.m^–2, resulting in a P/ ratio of 1.58. These results produced baseline information for the construction of a population-dynamics model and highlighted the potential of this species as an environmental quality-assessment bioindicator on sandy shores.Communicated by S.A. Poulet, Roscoff     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. II. Acartia omorii

Population dynamics and production of the calanoid copepod Acartia omorii Bradford were studied from November 1986 to November 1987 in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was present in the plankton from October to July (temperature range: 8.9 to 24.3°C), with peaks in February-March and June. During this period, nine generations could be detected, for which the mean population egg production rate and midstage abundance of each life stage older than naupliar Stage (N) II were determined to trace survival. The population suffered extremely high mortality during the early life stages: on average only 2.5% of the eggs produced recruited into NII. This large loss is probably concentrated within the egg stage, due to predation, including cannibalism, by omnivorous copepods, in addition to sinking loss of eggs in the water column. However, the mortality from NII to copepodite Stage (C) V was negligible, indicating low predation pressure by large carnivores. The biomass of A. omorii showed marked seasonal variations in parallel with numerical abundance. The instantaneous growth rate of each stage increased exponentially with increasing temperature. The integrated production rate of A. omorii from 7 November 1986 to 21 July 1987 was 749 mg Cm^-3 or 5.62 g Cm^-2     

The annual cycle of protozooplankton in the Kiel Bight

Protozooplankton (heterotrophic dinoflagellates and ciliates) composition and biomass was studied in a 20-m water column in the Kiel Bight on 44 occasions between January 1973 and April 1974. Both groups attained comparable biomass maxima during spring and autumn (0.3 to 0.7 g C m^-2 in the 20-m water column) and biomass levels were much lower in summer and lowest in winter. The spring protozooplankton maximum coincided with that of phytoplankton and during the rest of the year, protozooplankton stocks did not appear to be food limited as phytoplankton stocks were large throughout; many protozoans with ingested microplankton cells were observed, indicating that their potential food supply is not restricted to nanoplankton. Non-loricate organisms dominated biomass of the ciliates and tintinnids were of little importance. Tintinnids predominated in plankton samples concentrated by 20 m gauze indicating that most non-loricate ciliates, irrespective of size, were not retained. When phytoplankton sotcks were large (>3 g C m^-2) but those of metazooplankton small, as in spring and autumn, protozooplankton were the major herbivores with biomass levels comparable to those attained in summer by metazooplankton ( 0.5 g C m^-2). A highly significant negative correlation was found between protozooplankton and metazooplankton during the plankton growth season. Predation by the latter is thus an important factor regulating size of the protozooplankton population, although other factors also appear to be in operation. Loss rates of the pelagic system through sedimentation are highest in spring and autumn when protozooplankton dominate the grazing community and loss rates are much lower in summer when metazooplankton are the dominant herbivores. Apparently, the impact of protozooplankton grazing on the pelagic system is quite different to that of the metazooplankton.Publication No. 268 of the Joint Research Programme (SFB 95), Kiel University     

Population dynamics and growth of <Emphasis Type="Italic">Nassarius reticulatus </Emphasis>(Gastropoda: Nassariidae) in Rhosneigr (Anglesey,UK)

Seasonal changes in catch rate, growth and mortality of Nassarius reticulatus from an intertidal lagoon and a wave-exposed beach at Rhosneigr (Anglesey, North Wales, UK) are described. The number of N. reticulatus caught in baited traps from the lagoon was significantly higher (>125 individuals trap⁻¹) during the summer (>18°C), than at <12°C (<65 individuals trap⁻¹), and the numbers caught in the lagoon were an order of magnitude greater than on the beach, >13 individuals trap⁻¹ in July (>16°C), and <5 individuals trap⁻¹ between December and April (<9.5°C). Predictions of shell growth attained by N. reticulatus annually in the lagoon using graphical modal progression analysis (MPA) of length frequency data, were similar to the growth of marked and recaptured lagoon N. reticulatus. Predictions of shell growth using computerised length frequency distribution analysis (LFDA), however, did not reflect the growth as accurately as MPA. Modal progression analysis demonstrated that N. reticulatus from the lagoon achieved a higher asymptotic maximum shell length (L _∞) and a lower growth constant (K) than animals from the beach. Shell growth was seasonal with growth of the lagoon individuals slowing down towards the end of September and resuming in early April, about a month later than the beach individuals. Mortality of N. reticulatus was greater during the summer, and survival was lower in the lagoon than on the beach. Recruitment patterns were similar in the lagoon and on the beach, and MPA and LFDA predicted that larval N. reticulatus settled between late summer and early autumn, with juveniles (7–8.9 mm) appearing in the population the following year, between February and April. Growth of male and female N. reticulatus in the laboratory was similar and was temperature and size dependent. The different growth patterns between N. reticulatus from the two habitats, predicted using MPA, were maintained when individuals were reared under laboratory conditions for ∼6 months; N. reticulatus <21 mm from the beach grew faster than individuals from the lagoon, although N. reticulatus >21 mm from the lagoon grew faster and attained a larger length (26 mm) than individuals from the beach (24 mm). Low food availability did not affect N. reticulatus survival in the laboratory but significantly suppressed shell growth.     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Population dynamics of the tanaid Hargeria rapax (Crustacea: Peracarida) in a tidal marsh

The tanaidacean Hargeria rapax (Harger, 1879) was sampled along intertidal transects semi-monthly at one site and quarterly at two other sites in salt marshes on Sapelo Island, Georgia, USA, from July 1985 to July 1986. Tanaids were most abundant near the mean highwater line and became progressively less abundant at lower intertidal elevations. Population density was greatest in the winter (December to February) when there were >29 000 individuals/m² at one high intertidal station. Although reproductive individuals were present most of the year, peaks in reproductive activity occurred in autumn (late August to early November) and spring (early March to mid June). An increase in population density coincided with increased reproductive activity only in autumn. Tanaid cohorts produced in the spring and summer rarely survived beyond 6 to 8 wk, but those produced in the autumn overwintered and lived 22 to 26 wk. The sex ratio among mature individuals was 2.8:1 (females: males). Mature females ranged in size from 2.2 to 3.9 mm total length (TL) and mature males were 2.3 to 4.1 mm TL; there was no significant sexual difference (Student's t-test, P>0.05) in the mean TL of mature individuals. The mean (±SD) size of brooding females was 2.9±0.32 mm TL and the mean (±SD) nunber of offspring/brood was 8.3±4.99 young/female. The timing of tanaid reproduction together with the effects of predation by juvenile fish and crustaceans may account for most of the spatial and temporal patterns of tanaid abundance observed in this study. There was a significant linear relationship (P<0.001, r ²=0.54) between the growth rate (GR, mm/d) of individuals and average daily air temperature (°C) described by the equation: GR=0.00178 (°C)-0.00971. The potential annual contribution of tanaid production to higher trophic levels, estimated from knowledge of standing stocks, growth rates and fecundity, was 5.71, 0.91 and 0.46 g dry wt/m² for high, mid and low intertidal areas, respectively. The high intertidal marsh, which supports the largest and most persistent standing stock of H. rapax, provides a rich foraging area for aquatic predators at high tide and an important source of recruits from which tanaid populations at lower intertidal elevations are recolonized after periods of intense predation pressure.     

Animal community structure and energy budget calculations of a Daphnia magna (straus) population in relation to the rock pool environment

The fauna and environmental conditions in a freshwater rock pool ecosystem were followed by a weekly quantitative sampling program from April to August, 1974. The rock pool, situated on a small island in the northern Baltic Sea, was heavily eutrophicated by droppings from the surrounding colony of sea birds. The intermittent flushing of the pool with rainwater and the input of bird droppings, as well as the biological activity, contributed to the large seasonal variations in nutrients and organic matter that were observed. A dense algal bloom of flagellates occurred in April but vanished when the animal population started to increase in the middle of May. During the rest of the summer, most of the photosynthetic pigments were found in the bottom sediment, mostly as degraded phaeo-pigments due to intensive grazing by the animals in the pool. Very few animal taxa were found and the phyllopod Daphnia magna (Straus) dominated throughout the whole sampling period. D. magna contributed to more than 50% of the total biomass, except in late July when chironomid larvae were most abundant. The total biomass in the pool increased from about 15 mg (dry weight) l^?1 in May to a maximum of about 60 mg l^?1 at the end of June. There were few carnivores in the system, except during the spring when the water bug Deronectes griseostriatus (de Geer) was common.Analyses of size, age and sex structure and calculations of birth and death rates of the D. magna population showed large seasonal variations, correlated with the volume fluctuations and flushings of the pool which stimulated both the algal production and the growth and reproduction of Daphnia. Data from the extensive literature that exists on D. magna and other species of the same genus were used, together with field and experimental data from the rock pool population, in a numerical model describing the energy budget of this species. The model describes variations in weight-specific growth, reproduction, moulting, feeding and respiration rates in relation to temperatures and food concentrations. Energy budget relationships that maximise the chances for survival and utilisation of the available energy for a population exposed to varying food concentrations are predicted by the model. The energy budget model was also used to estimate the secondary production of the rock pool population of D. magna. The total production between April and August was 203 mg (dry weight) l^?1 and the average production per biomass ratio was 0.094 day^?1. The average net production efficiency was 42%, very close to other values reported for D. magna from other biotopes. The relative importance of different factors controlling the production was also analysed with the model. Production per biomass ratio was calculated, assuming a constant temperature and/or food concentration for the whole sampling period. The varying food concentrations in the rock pool had the most pronounced influence, greater than that of temperature and size structure variations in the population.     

Seasonal variations in distribution and population structure of Microcalanus pygmaeus and Ctenocalanus citer (Copepoda: Calanoida) in the eastern Weddell Sea,Antarctica

The abundance, vertical distribution and population structure of two important small calanoid copepod species, Microcalanus pygmaeus (G. O. Sars) and Ctenocalanus citer Heron and Bowman, were studied in the eastern Weddell Sea in summer (January/February 1985), in late winter/early spring (October/November 1986) and in autumn (April/May 1992). The population of Microcalanus pygmaeus consisted mainly of copepodite stages CII and CIII in late winter/early spring and were concentrated between 500 and 200 m depth. In summer, stage CIV was the modal stage and the bulk of the population had ascended above 300 m. In autumn the population structure was bimodal with CI and CV dominating. Most of the population was concentrated between 300 and 200 m. In all investigation periods M. pygmaeus had their maximal concentrations in the thermo-pycnocline. The developmental stages CIII to CV of Ctenocalanus citer formed the bulk of the population in late winter/early spring. In October all developmental stages had their main distribution between 500 and 200 m, except females, which were concentrated in the upper 50 m. In November most of the population occurred between 200 and 50 m. The summer population was concentrated in the upper 50 m, and numbers increased dramatically as the new cohort hatched. Copepodite stages CII and CIII dominated the population at the end of January, while CIV dominated 2 wk later. In autumn, CV was the modal stage. The majority of the population was concentrated in the upper 100 m, but there was an increase in abundance below 300 m compared to summer. Age structure changed with depth with a younger surface population and an older one in deeper water layers. The seasonal change in number of M. pygmaeus is much smaller than that of C. citer; the summer:winter:autumn ratio of the former being about one, whereas the winter:summer/autumn of the latter was about nine. Early copepodite stages and adults of M. pygmaeus occurred throughout all investigation periods. The large proportion of early copepodite stages in April and in mid-October suggests autumn and early to midwinter breeding. Apparently, M. pygmaeus may reproduce and grow year-round or perhaps has a 2-yr life-cycle. In contrast, the dramatic increase in abundance of early copepodite stages of C. citer in summer suggests springtime reproduction.     

Seasonal occurrence of the ctenophore Pleurobrachia pileus in the western Dutch Wadden Sea

Due to the short residence time of the water in the western Dutch Wadden Sea the ctenophore Pleurobrachia pileus has to be considered as an allochthonous species, which enters the estuary passively from the North Sea. The seasonal occurrence of this species in the Dutch Wadden Sea is a reflection of its abundance in the coatal zone of the North Sea and is characterized by a pronounced spring peak of up to 17000 individuals per 1000 m³ and a lower autumn peak with numbers up to 400 individuals per 1000 m³. Similar to other areas, predation by Beroë gracilis, another ctenophore species, may be a factor in controlling the P. pileus population in this region, although B. gracilis abundance is low. P. pileus is only important as a predator for a short time, i.e. the month of May, when it reaches maximum biomass values of up to 12 00 mgC per 1000 m³. Both the estimated potential consumption rates in comparison with the available food as well as the observed suboptimal growth rate of the population suggest that in the coastal zone of the North Sea area food conditions may not be optimal for P. pileus.     

Interlinked temporal changes in environmental conditions, chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal sandflat (Seto Inland Sea, Japan)

Five field surveys were conducted in an estuarine intertidal sandflat of the Seto Inland Sea (Japan) between April 1994 and April 1995. Chlorophyll a, pheopigments, total organic carbon and acid-volatile sulphides (AVS) of surface and subsurface sediments, and macrofaunal assemblages were investigated in parallel at 15 stations. Monthly hydrological data of low-tide creek water adjacent to the flat were used as a complementary environmental characterisation of the study area. Strong temporal changes were found among sampling dates, most remarkably in autumn with a major increase of algal detritus and AVS, a sharp reduction in macrofaunal abundances and species richness, and a massive mortality of the clam Ruditapes philippinarum. This dystrophic event was preceded by a photoautotrophic and hypertrophic spring–summer characterized by abundant fresh (i.e., living) algal material, including microphytobenthos and macroalgae (Ulva sp.). In summer, abundant macrofaunal assemblages reached the highest biomass values (455 g wet weight m⁻² or 60.6 g ash free dry weight m⁻²), with a major contribution of filter-feeding bivalves Musculista senhousia and R. philippinarum. These are among the highest values reported in the literature for sedimentary shores. From autumn, there was a progressive recolonisation of macrofauna, initiated by few opportunistic polychaetes (e.g., Cirriformia tentaculata and Polydora sp.), apparently promoting a fast sediment recovery in winter, and followed by new bivalve recruits in the next spring. This study provides the first evidence of significant and interlinked within-year changes in chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal flat at a small spatial scale (i.e., tens of meters). This demonstrates the high temporal variability of species–environment relations in these systems and a close relationship in seasonally driven trophodynamic processes among primary producers and benthic consumers. We conclude that a thorough parallel evaluation of the temporal changes in chemical characteristics of sediments should be taken into account in assessing the year-round distribution and changes of intertidal macrofauna, particularly in eutrophic, estuarine intertidal flats.An erratum to this article can be found at     

Population biology of hyperbenthic crustaceans in a cold water environment (Conception Bay,Newfoundland). I.<Emphasis Type="Italic"> Mysis mixta</Emphasis> (Mysidacea)

Seasonal population dynamics of Mysis mixta Lilljeborg were studied from December 1998 to November 2000 at a 240 m deep site in Conception Bay, Newfoundland. At this depth, temperature was <0°C and salinity between 32.0 and 34.0 psu year-round. The spring phytoplankton bloom began in early or late March and reached a maximum in late April to mid-May. M. mixta exhibited a highly synchronised life cycle, with spawning and mating occurring in October to November, embryos brooded for ~5 months, and juveniles released during spring bloom sedimentation in April and May. Females were semelparous and died at age 2.5 years, following release of juveniles in spring, whereas the majority of mature males died at age 2 years, following mating in November. The biennial life cycle of this population resulted in the presence of two cohorts in the hyperbenthos at any given time. Variation in density and biomass was low among cohorts but high within cohorts, the latter probably due to the high motility of mysids. Densities in 1999 and 2000 were 242±379 and 544±987 ind. per 100 m³ (mean±SD), respectively. Although growth rates were similar between years, rates measured from changes in dry mass differed both seasonally and among life-history stages (range from –4 to 7 mg month^–1). Annual secondary production was estimated at 29–73 mg C m^–2 in 1999 and 53–205 mg C m^–2 in 2000. The annual P/B ratios were 1.62 and 1.19 in 1999 and 2000, respectively.Communicated by J.P. Grassle, New Brunswick     

Changes in yolk total proteins and lipid components and embryonic growth rates during lobster (<Emphasis Type="Italic">Homarus americanus</Emphasis>) egg development under a simulated seasonal temperature cycle

From August 2000 to June 2001, seven egg-carrying female lobster (Homarus americanus) from the Îles de la Madeleine population (Gulf of St. Lawrence, Canada) were held under a simulated seasonal temperature cycle to monitor egg development from extrusion to hatching. For the first time, changes in the yolk components (total lipids and major lipid classes, total proteins) and embryo growth of single eggs were monitored separately over the entire development period. Under the controlled temperature conditions, egg development proceeded in three phases. (1) Autumn, from extrusion to early December, was marked by a rapid increase in the Perkinss eye index and rapid declines in yolk total proteins and triacylglycerols (TAG). Embryo daily growth rate was estimated between 1 and 2 µg proteins day^–1. (2) Winter, from late December to early April (temperature stable at ca. 1°C) was characterized by a stationary phase in the evolution of the eye index and yolk lipid use, and embryo growth slowed significantly. (3) Spring, from late April to hatching in June was the period with the most rapid changes in yolk TAG and embryo growth rates >6 µg proteins day^–1 were recorded. Almost 65% of the live biomass (total proteins) of the hatching larvae was accumulated during the last few weeks of development. An index of embryo growth efficiency was estimated as the slope of the relationship between embryo total proteins and yolk TAG during egg development. A relationship was found between the initial mean egg dry weight and the embryo growth efficiency index suggesting that under the same experimental conditions bigger eggs used yolk lipids more efficiently and sustained faster embryonic growth than smaller eggs. The relationship may also explain why larger larvae originate from larger eggs.Communicated by R.J.Thompson, St Johns     

The effects of temperature on the growth of juvenile scleractinian corals

Tropical reef corals are well known for their sensitivity to rising temperature, yet surprisingly little is known of the mechanisms through which temperature acts on intact coral colonies. One such mechanism recently has been suggested by the association between the growth of juvenile corals and seawater temperature in the Caribbean, which suggests that temperature causes a transition between isometric and allometric growth scaling in warmer versus cooler years, respectively (Edmunds in Proc R Soc B 273:2275–2281, 2006). Here, this correlative association is tested experimentally for a cause-and-effect relationship. During April and May 2006, juvenile colonies (8–35 mm diameter) of massive Porites spp. from Moorea, French Polynesia, were incubated at warm (27.8°C) and cool (25.7°C) temperatures for 15 days, and their response assessed through the scaling of growth (change in weight) with colony size. The results reveal that the scaling of colony-specific growth (mg colony⁻¹ day⁻¹) was unaffected by temperature, although growth absolutely was greater at the cool compared to the warm temperature, regardless of colony size. This outcome was caused by contrasting scaling relationships for area-specific growth (mg cm⁻² day⁻¹) that were negatively allometric under warm conditions, but independent of size under cool conditions. In April 2007, a 22 days field experiment confirmed that the scaling of area-specific growth in juvenile Porites spp. is negatively allometric at a warm temperature of 29.5°C. Based on strong allometry for tissue thickness, biomass, and Symbiodinium density in freshly collected Porites spp., it is hypothesized that the temperature-dependency of growth scaling in these small corals is mediated by the interaction of temperature with biomass.     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. III.Paracalanus sp.

Population dynamics and production of the calanoid copepodParacalanus sp. were studied from November 1986 to November 1987 in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large abundance peak in June/July and small peaks in September/October and November/December. During a year of investigation, 15 generations Gould be detected. For each generation, the mean population egg production rate and the mean daily midstage abundance front NIII to CV were determined to obtain a survival curve from egg to CV. The mortality was extremely high during the early life stages: on average only 7.1% of the eggs produced might survive into NIII. This high mortality might be caused by predation by sympatric omnivorous copepods, in addition to sinking loss of eggs from the waten column. The biomass ofParacalanus sp. showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous growth rate of each developmental stage increased exponentially with temperature up to 20 °C, above which the rate was constant. The annual integrated production rate was 734 mg C m^–3 yr^–1 or 5.5 g C m^–2 yr^–1.     

Temporal variation of Tubularia indivisa (Cnidaria, Tubulariidae) and associated epizoites on artificial habitat communities in the North Sea

We have analyzed the composition, diversity, density and biomass of a temporal series of samples taken in a Tubularia indivisa community, which dominates a shipwreck in the North Sea waters (N 51°23′,730–E 02°29′,790, 17 nautical miles from the coast, 30 m depth). This shipwreck has structures emerging up to 8 m above the seabed. Water temperature ranged from 4.2°C in March to 20.3°C in August. Salinity showed few variations around 33.9 psu. Bottom tidal currents followed a semi-diurnal cycle and were preferentially NE oriented with 84% of them in the range 0.25–0.75 m s⁻¹. The mean value for total suspended matter was 6.2 mg l⁻¹ with large variations on a monthly scale. The species richness of samples varied from 15 in October to 42 in August with a mean value of 33 species. Diversity indices were higher during autumn and winter because of the strong dominance of a few crustacean species during the warmer months. The total density of individuals ranged from 6,500 ind m⁻² in October to 445,800 ind m⁻² in July, most of these individuals belonging to the amphipod species Jassa herdmani. The biomass of the T. indivisa community varied from 9 g AFDW m⁻² in October to 1,106 g AFDW m⁻² in July, with T. indivisa itself constituting between 59 and 82% of the total biomass. The biomass of T. indivisa was positively correlated with species richness and with the density of 23% of the species identified on this community, suggesting that T. indivisa plays an important structural role in this habitat. This was further confirmed by the number of species associated with T. indivisa which was generally superior to 55% of the sorted species. Multivariate analysis indicated strong differences between spring/summer−autumn/winter assemblages mostly but not solely due to the abundance patterns of species. These findings support the conclusion that shipwrecks in Belgian waters allow the development of assemblages dominated by a high biomass of T. indivisa which in turn provides shelter for high densities and biomass of epizoites. These assemblages will further show large monthly variations in densities and composition due to large variation in T. indivisa biomass under an apparent repetitive annual cycle. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A comparison of macrofaunal and meiofaunal distribution and standing stock across a rocky shore,with an estimate of their productivities

In 1984 and 1985 algal, macrofaunal and meiofaunal standing stocks were estimated on a exposed rocky shore along the west coast of False Bay, South Africa, using comparable, area-based sampling techniques. The shore supported a rich growth of algae, particularly in summer, when a maximum standing crop of 403 g m^-2 was recorded in the low shore. In winter, the largest component of macrofaunal biomass comprised the filter-feeding barnacle Tetraclita serrata, which attained 75 g m^-2 in the middle balanoid; but as a result of late recruitment and high mortality of this species, the summer shore was dominated by herbivorous gastropods, particularly Patella cochlear, which attained a maximum biomass of 66 g m^-2 on the low shore. Meiofaunal numbers and biomass were closely correlated to the distribution of algal turfs and associated trapped sediments. Numerically, the most important components of the meiofauna were nematodes and copepods, while the biomass was more evenly shared among foraminifera, minute gastropods, copepods and insect larvae. Numbers and biomass peaked in the lower balanoid during winter (1.9×10⁶ individuals, or 8.5 g m^-2). Macrofauna:meiofauna numbers and biomass ratios are presented for each zone and the distribution patterns discussed in relation to the conditons in each. Numerically, meiofauna exceed macrofauna by an overall ratio of 1:391, with values ranging from 1:556 in the lower balanoid to 1:18 in the Littorina zone. Macrofaunal biomass exceeds that of meiofauna in all zones by an overall ratio of 10:1, but values range from 2.1:1 in the upper balanoid to 48:1 in the middle balanoid. By incorporating turnover ratios extrapolated from the literature, mean annual productivity ratios have been calculated. These indicate that macrofauna account for 75% of total secondary production and meiofauna for 25%. Failure to incorporate meiofauna in analyses of energy flow on rocky shore ecosystems would thus lead to considerable errors. The possible trophic role of meiofauna in such systems is discussed.